First published online June 16, 2005
Journal of Experimental Biology 208, 2549-2554 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01665
Emperor penguins adjust swim speed according to the above-water height of ice holes through which they exit
Katsufumi Sato1,*,
Paul J. Ponganis2,
Yoshiaki Habara3 and
Yasuhiko Naito4
1 International Coastal Research Center, Ocean Research Institute,
University of Tokyo, 2-106-1 Akahama, Otsuchi, Iwate 028-1102,
Japan
2 Center for Marine Biotechnology and Biomedicine, Scripps Institution of
Oceanography, University of California San Diego, La Jolla, CA 92093-0204,
USA
3 Laboratory of Physiology, Department of Biomedical Sciences, Graduate
School of Veterinary Medicine, Hokkaido University, North 18, West 9, Sapporo
060-0818, Japan
4 Biologging Institute, 2-31-10, Rex Yushima 301, Yushima Bunkyo, Tokyo
113-0034, Japan

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Fig. 1. Schematic diagram of exit options for the penguins. As shown on the left,
the bird could climb onto the ice if the center of gravity reached the height
of the ice around the hole, or the exit speed was sufficient to project the
bird onto the ice (the right-hand bird).
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Fig. 2. Typical profiles for last 20 s before exit through the ice hole in 6th
deployment by bird no. 314. The closed circle indicates the maximum swim speed
before exit. Horizontal bars indicate strokes based on acceleration.
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Fig. 3. Relationship between above-water height and the maximum speed before exit.
Observed data (386 exits in 13 deployments by nine birds are pooled) are
represented by closed circles (successful exit) and crosses (failed exit). The
solid curve is the theoretical line for initial vertical speed to reach the
height, assuming a take-off angle ( ) of 90° (see text for details).
If penguins adopt non-vertical body angles, higher swim speeds were needed to
obtain the necessary vertical speeds for the height, which are represented by
dotted curves for various take-off angles.
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Fig. 4. Box plots for the comparison in the maximum speed before exit between lower
(white bars) and higher (gray bars) holes for each deployment. The differences
in the above-water height of holes were less than 10 cm in four deployments
(from 7th to 10th). *P<0.05;
**P<0.01; ***P<0.001; two-tailed
Mann-Whitney U tests; n.s., not significant.
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Fig. 5. An example of the last 12 s before exit through the ice hole in the 8th
deployment by bird no. 429. The closed circle indicates the maximum swim speed
before exit. Horizontal bars indicate strokes based on acceleration.
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© The Company of Biologists Ltd 2005